ACTA PALAEONTOLOGICA ROMANIAE (2018) V. 14 (1), P. 31-45
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1Geological Institute of Romania - 1st, Caransebeş Street, Ro-012271, Bucharest, Romania. E-mail: [email protected] 31 2Istanbul University Forestry Faculty, Department of Forest Botany, 34473 Bahceköy-Sarıyer, İstanbul / Turkey.
*Corresponding author: [email protected]
NEOGENE PALMOXYLON FROM TURKEY
Stănilă Iamandei1, Eugenia Iamandei1 & Ünal Akkemik2*
Received: 13 May 2018 / Accepted: 26 July 2018 / Published online: 2 August 2018
Abstract Tertiary fossil wood identifications of Turkey revealed many different gymnosperm and angiosperm genera
and species. Within this lignoflora some Palmoxylon types were identified for the first time as Palmoxylon coryph-
oides Ambwani et Mehrotra, Palmoxylon sp. cf. Trachycarpus H. Wendl., and Palmoxylon sp. cf. Borassus L. The
fossil palm-wood was collected from the sites in Seben and Gökçeada from early Miocene deposits, and from the site
in Erikli from middle-late Miocene deposits. The presence of the palm trees in the Miocene Flora of the Anatolian
and Aegean regions indicates a lowland riparian palaeoenvironment and warm-humid or subtropical palaeoclimate.
Keywords: Palmoxylon, fossil-species, Seben, Gökçeada, Erikli.
INTRODUCTION
In the recent decades, our information on the fossil lig-
noflora of Turkey has increased. The new identifications
of petrified wood in Turkey showed a rich woody flora
from the late Oligocene to the late Miocene. Some of the
identified genera such as Sequoioxylon (Özgüven-Ertan,
1971, Kayacik et al., 1995; Akkemik et al., 2005; Sakınç
et al., 2007, Akkemik & Sakinç, 2013, Akkemik et al.,
2009), Glyptostroboxlon (Akkemik et al., 2017), Podo-
carpoxylon (Sakınç et al., 2007), Engelhardioxylon
(Sakınç et al., 2007) are taxa completely extinct from
Turkey, while other fossil forms (e.g. Sayadi, 1973; Sel-
meier, 1990; Dernbach et al., 1996; Aytuğ & Şanli, 1974;
Eroskay & Aytug, 1982; Şanli, 1982; Akkemik et al.,
2016 and Bayam et al., 2018) still have modern corre-
spondents. Regarding with Palmoxylon, many studies
were published in the world (e.g. Schenk, 1882; Mahaba-
lé, 1958; Greguss, 1959, 1969; Kaul, 1960; Tomlinson,
1961, 1990; Prakash, 1962; Grambast, 1962, 1964;
Trivedi & Verma, 1970; Roy & Ghosh, 1980; Privé-Gill
& Pelletier, 1981; Ambwani & Mehrotra, 1990; Gott-
wald, 1992; Nambudiri & Tidwell, 1998; EL-Saadawi et
al., 2004; Kahlert et al., 2005; Sakala, 2004; Dransfield et
al., 2008; Thomas, 2011a,b; Thomas & De Franceschi,
2012, 2013; Kamal-EL-Din et al., 2013; Prasad et al.,
2013; Nour-EL-Deen et al., 2017). Otherwise an extend-
ed list o References on Palmoxylon is published in the
site CiteULike by Millevacs in: (Accessed 05.05.2018).
Within these studies Kamal-EL-Din et al. (2013) stated
that Egypt is the richest African country having fossil-
species of Palmoxylon with 16 species, after describing
four Palmoxylon species, P. deccanense Sahni, P. ed-
wardsi Sahni, P. geometricum Sahni, P. prismaticum
Sahni, P. pondicherriense Sahni, P. pyriforme Sahni and
P. sagari Sahni.
Thomas & De Franceschi (2013) made a valuable revi-
sion of the palm xylotomy, which is very useful for mod-
ern palm identification by the study of the palm stem mi-
croscopic structure and also useful for the fossil palms, as
species of Palmoxylon. Descriptions were based mainly
the general organization of the transverse section and the
structure of the fibrovascular bundles as well as the
ground parenchyma, taking the Coryphoideae as case-
study.
As for Anatolian region, the palaeobotanical studies men-
tioned above revealed that the climate in the early Mio-
cene was warmer, probably subtropical and much humid
than the present climate. Palm trees growth even today
under this kind of climate in Seben (Akkemik et al.,
2016), European Part of Turkey and in the Island of
Gökçeada in Aegean Sea (Güngör et al., 2018), but their
distribution area was much wider through Turkey during
Neogene. Under this discussion, the purpose of the pre-
sent paper is to describe some fossil forms based on the
study of their petrified remains.
MATERIAL AND METHODS
We had in study four samples that sowed the typical palm
structure and were considered as fossil forms of Palmoxy-
lon. They were coded as HOC25 (Akkemik et al., 2016),
GOK01 and GOK269 (Güngör et al., 2018), and ERI01.
They have been collected from Bolu-Seben-Hoçaş Fossil
Site (HOC25), Edirne-Erikli (ERI01) and Gökçeada
(GOK01 and GOK269) (Fig. 1). The samples were in-situ
position in Bolu-Seben-Hoçaş Fossil Site (Fig. 2) and
dispersed in the others (Fig. 3).
According to Akkemik et al. (2016), the age of fossil site
in Seben-Hoçaş was 18.2±0.8 from the basalt samples.
Similar ages were given by Keller et al. (1992), Toprak et
al. (1996), and Wilson et al. (1997) including the ages of
17.6-24.8 Ma for volcanic activity in Galatean Volcanic
Province. So, the age of Seben-Hoçaş Fossil Site is the
early Miocene.
Güngör et al. (2018) specified that the geological age of
Gökçeada as also early Miocene, bearing a similar wood
flora with Lesbos Island (Süss & Velitzelos, 1997).
The fossil wood material (ERI01) in Erikli was found on
the shore of Aegean Sea. Sakinç et al. (1999) explained
geological background of Erikli, and its age is most prob-
ably middle-late Miocene.
Stănilă Iamandei, Eugenia Iamandei & Ünal Akkemik
32
Fig. 1 The locations of the sampled materials.
Fig. 2 In-situ trunk of Palmoxylon in Seben-Hoçaş Fossil Site.
Fig. 3 Dispersed wood trunk fragments including one of Palmoxylon in Gökçeada Fossil Site.
Neogene Palmoxylon from Turkey
33
For wood identification, we realized thin sections from
the samples of fossil wood along the three standard
planes: transverse section (TS), or cross-section, radial
longitudinal section (RLS) or simply, radial section and
tangential longitudinal section (TLS) or simply tangential
section. These standard planes of cutting in a palm stem
are also valid since there is an oriented structure in the
external and intermediary part, only in the central part of
the stem is less important (see Mahabalé, 1958; Kaul,
1960; Greguss, 1959, 1968; Tomlinson, 1961)
The samples and sections were stored at the "Laboratory
of Tree-ring Researches and Wood Anatomy" in Forest
Botany Department of Faculty of Forestry, Istanbul Uni-
versity.
The first study was made on a biological microscope
(LEICA DM2500 Light Microscope) and the micro-
photos were realized with a LEICA DSC295 type of
camera.
The description and identification of the specimens was
performed in the Geological Institute of Romania, in Bu-
charest. In identification, many references including fos-
sil Palmoxylon species were used (e.g. Schenk, 1882;
Mahabalé, 1958; Greguss, 1959, 1968, 1969; Kaul, 1960;
Tomlinson, 1961, 1990; Prakash, 1962; Grambast, 1962,
1964; Trivedi & Verma, 1970; Roy & Ghosh, 1980;
Privé-Gill & Pelletier, 1981; Ambwani & Mehrotra,
1990; Gottwald, 1992; Nambudiri & Tidwell, 1998; EL-
Saadawi et al., 2004; Kahlert et al., 2005; Sakala, 2004;
Dransfield et al., 2008; Thomas, 2011a, b; Thomas & De
Franceschi, 2012, 2013; Kamal-EL-Din et al., 2013; Pra-
sad et al. 2013; Nour-El-Deen et al., 2017).
SYSTEMATIC PALAEOBOTANY
Order Arecales Bromhead 1840 (in Reveal, 2004)
Family Arecaceae Berchtold et Presl, 1820 (nom. cons)
Genus Palmoxylon Schenk, 1882
Palmoxylon coryphoides Ambwani & Mehrotra, 1990
Fig. 4, photos a-i. Fig. 5, photos a-i.
Material: GOK01 and GOK259.
Locality: Gökçeada.
Formation: Kesmekaya Volcanics.
Age: Early Miocene.
Origin on the stem remains: Coming from unknown
portion of the trunk.
Storage: This material is now kept under index GOK01
and GOK269 within the "Collection of fossil woods" in
the Forest Botany Department, Forestry Faculty, Istanbul
University.
Microscopic description: Theoretically the central cyl-
inder, in cross section, is formed from three distinct parts:
dermal, subdermal and central zones (see Mahabalé, 1958
and Kaul, 1960 quoting von Mohl, 1850) or external,
intermediary and central (in euro-american school, see
Tomlinson, 1961), or, respectively, subcortical zone - as a
periferal sclerotic zone, transitional zone and central zone
as was lastly stated by Thomas & De Franceschi (2013).
For description, the shape, the size, the density of the
fibrovascular bundles, their fibrous part and vessels, the
quantity of parenchyma, the abundance and the arrange-
ment of parenchymatous cells starch grained along to the
fibrovascular bundles must be considered and also the
phytoliths.
The studied specimens show in cross-section the central
zone, but also at least a part of the transitional zone (es-
pecially the specimen GOK259), where fibro-vascular
bundles of open collateral type appear, usually oriented
with the vascular part to the centre of the stem and the
fibrous part to outside, and having 2-4 large metaxylem
vessels round to oval, and typical reniform sclerenchyma
caps well developed, floating in the ground parenchyma
tissue, often touching each other. The size of fvb is varia-
ble 400-500/360-475 µm (mean values: 421.25 / 403.75
μm, the phloematic (=anterior) sclerenchyma cap has
r/tg.d = 116 / 372 μm, f / v ratio is 0.35 / 1 and the fre-
quency, or density, is of 491.2 bundles/cm2.
The phloem appears protected under the median sinus of
the sclerenchyma cap, a single island - as an undivided
sieve plate, rarely apparently divided.
The metaxylem appear as 2-4(-6-8) vessels either oval or
elliptic or, less round in cross-section or, sometimes, par-
titioned by the terminal inclined wall. They are usually
large and thick-walled vessels, having in cross section 70-
162.5 / 45-107.5 μm (mean values 107.7 / 65 μm) in di-
ameters and 8-10 µm the simple wall. The size of meta-
xylem vessels is of 47 / 44.55 μm, and thick-walled, of 7-
10 µm the simple wall. Vertically the walls of metaxylem
vessels have scalariform pitting and annular thickenings
sometimes ramified and anastomosed. The terminal wall
is very inclined and bears a scalariform perforation with
8-18 thick bars and, sometimes, only simple perforation.
The protoxylem appear as unequally smaller vessels
grouped beneath the metaxylem vessels or partially
pushed between them, possibly because the sustained
growing of the parenchyma from around. They have
d=25/22.9 μm, and are relatively thick-walled: 3-5 μm in
the simple wall.
The intrafascicular (or paravascular) parenchyma is con-
stituted of uniform, relatively thin-walled (5-7 μm the
simple wall), rounded-polyhedral cells which appear also
rounded in cross-section and small-sized (15.5/5 - 25/10
μm). Adjacent to the metaxylem vessels appear in 1-2
rows regularly arranged, similar to a tabular parenchyma.
The marginal 1-2 rows of sclerified paravascular paren-
chyma cells constitute a second sclerenchyma cap pro-
tecting the vascular zone also named ventral fibrous cap
adjacent to the xylem.
The ground tissue (or the interfascicular parenchyma) is
constituted by non-oriented rounded-polyhedral cells
unequally sized, very emaciated or lobed cells, as they
appear in cross section, usually thin-walled, and with
possible starch-content. It represents the ground paren-
chyma with sustained growing of Corypha type. All
around the fibrovascular bundle only tabular parenchyma
can be seen, in 1-2 compact rows arranged. Fibrous bun-
dles weren't observed.
The central zone in both the specimens studied is of Cor-
ypha type (von Mohl, in Thomas & De Franceschi, 2011)
where the round to oval fibrovascular bundles are varia-
bly oriented, having 2-4(-6-8) metaxylem vessels and not
too developed sclerenchyma caps of Reniforma to Lunar-
ia type, all floating in the ground parenchymal tissue
along with small fibrous bundles, more numerous in
Stănilă Iamandei, Eugenia Iamandei & Ünal Akkemik
34
Fig. 4 Palmoxylon coryphoides Ambwani & Mehrotra, 1990. (GOK01). a-c Cross-section. Fibrovascular bundles (fvb)
variably oriented in central zone; parenchymal ground tissue; fibrous bundles; c Detailed fvb-s displaying metaxylem and
protoxylem vessels, phloem, reniform sclerenchyma cap, stegmata, and rounded by tabular parenchyma. Also ground
tissue parenchymal cells and fibrous bundles can be seen. d-i Longitudinal sections. Metaxylem and protoxylem vessels
with annular thickenings, scalariform pitting, and with scalariform perforated inclined plates. Also, long rows of stegmata
can be seen. g, i Metaxylem and protoxylem vessels with annular bifurcated thickenings, and with scalariform perforated
plate. h Metaxylem vessel with scalariform pitting.
Neogene Palmoxylon from Turkey
35
Fig. 5 Palmoxylon coryphoides Ambwani & Mehrotra, 1990. (GOK259). a-c Cross-section. a Transitional zone with fvb
with reniform caps; parenchymal ground tissue; b Variably oriented fvb in central zone; parenchymal ground tissue; fi-
brous bundles; c Detailed fvb coupled with small fvb-s displaying metaxylem and protoxylem vessels, phloem, reniform
sclerenchyma cap, stegmata, and tabular parenchyma. Also ground tissue parenchymal cells and fibrous bundles can be
seen. d-g Longitudinal sections. Metaxylem and protoxylem vessels with annular or helical thickenings, scalariform pit-
ting, and with scalariform perforated inclined plates. Fibrous sheaths of fvb marked by long rows of stegmata (d, f). g-i
Metaxylem and protoxylem vessels with annular and helical thickenings and with simple perforation (i).
Stănilă Iamandei, Eugenia Iamandei & Ünal Akkemik
36
GOK01 specimen. Under the sclerenchyma cap a large
median sinus is present, auricular sinuses were not ob-
served. The sclerenchyma cap (also named anterior or
dorsal cap, or phloematic fibrous sheath, or fibrous part
adjacent to phloem) is constituted from vertically elon-
gated fibrous cells with polygonal rounded cross-section,
thick-walled and with rounded to point-like lumina. The
size of the fibrovascular bundles is variable: 400-
500/360-475 µm (mean values: 421.25/403.75 μm, the
phloematic (=anterior) sclerenchyma cap has
r/tg.d=116/372 μm, f/v ratio is 0.35/1 and the frequency
(or density) is of 214 bundles/cm2. Signs of a centrifugal
differentiation of the fibrous part seem to be absent.
The phloem, appear protected under the median sinus of
the sclerenchyma cap, in a single island - as an undivided
sieve plate, rarely apparently divided.
The metaxylem appear as 2-4(-6-8) vessels either oval or
elliptic, or less, round in cross-section or, sometimes par-
titioned by the terminal wall. They are usually large and
thick-walled vessels (7-8-10 µm the simple wall), having,
in cross section, 70-162.5/45-107.5 µm (mean 107.7/65
µm) in diameters (smaller in specimen GOK259:
47.03/44.55). Vertically the walls of the metaxylem ves-
sels have scalariform pitting and annular thickenings
sometimes ramified. The terminal wall is very inclined
and bears a scalariform perforation with 8-18 thick bars
(fewer in the specimen GOK259: 3-5-8 bars) or even
simple perforation.
The protoxylem is represented by 3-12 small round to
oval vessels of 10-36 µm in diameters, more numerous in
the specimen GOK259 (up to 30), and relatively thick
walled (2-5 µm the simple wall). Vertically the protoxy-
lem vessels present annular thickenings sometimes rami-
fied, and short scalariform perforations.
The intrafascicular (or paravascular) parenchyma is con-
stituted of uniform, thin-walled (1-2 µm the double wall),
rounded-polyhedral cells which appear polygonal-
rounded in cross-section, small-sized cells (round of 6-14
µm or elliptic of 12.5-25/5-15 µm). Adjacent to the meta-
xylem vessels appear in 1-2 regular rows of flattened
cells arranged, like a tabular parenchyma. All the re-
mained space is filled with compact non-oriented paren-
chyma cells. The marginal 1-3 rows of paravascular pa-
renchyma is constituted by highly sclerified cells, thicker
walled (3-6 µm the double wall), so forming a second
sclerenchyma cap, which protect the vascular zone, also
named ventral cap or fibrous part adjacent to the xylem.
The ground tissue or the interfascicular parenchyma is
constituted by non-oriented very emaciated big cells as
lobed cells, as they appear in cross section, usually thin-
walled, and with possible starch content. It represents the
ground parenchyma with sustained growing of Corypha
type, the cells compressing each other. All around the
fibrovascular bundle only tabular parenchyma can be
seen, in 1-2 compact rows arranged.
The fibrous bundles are more numerous in specimen
GOK01, floating in the ground parenchyma among the
fibrovascular bundles. They have 15-19-25 fibers, or
more, with polygonal rounded cross-section, very thick
walls and point-like lumina. Often the fibers bear phyto-
liths as spherical stegmata, slightly spinulose, partially
sunken in the basal wall of silica-cell wall. They are visi-
ble even in cross section, and appear in long rows in the
longitudinal sections.
The phytoliths as spherical stegmata also appear, vertical-
ly, on the fibers of fibrous part of the fibrovascular bun-
dles, numerous and in long rows arranged, covering large
surfaces on the fibrous part of the bundle, also visible in
cross-section around caps. Those spherical stegmata seem
to bear rounded spines, and are partially sunken in the
basal wall of the silica-cell. It seems that these stegmata
had from the beginning a definitive size, since it seems
that in time they didn’t grew more. Also, they seem to
connect the fibers with one another, thus enhancing the
solidity of the structure.
Affinities and discussions: In cross section, the studied
material presents a monocotyledonous fascicular struc-
ture, typical for the stem of palm tree, a member of the
Family Arecaceae Berchtold et Presl, 1820 (nom.cons)
also found in the scientific literature as Family Palmae
Jussieu, 1789 (nom. cons. et nom. alt.) or as Family Are-
caceae Schultz-Schultzenstein, 1832 (nom.cons.). The
fossil correspondent, Palmoxylon is a genus name that
generally defines “wood of Palm”, in conformity with the
original diagnosis of Schenk (1882). However, taking
into account that other organs of the palm-plant are con-
nected with the stem (rootlets, leaf axes, petioles, fruits,
flowers, pollen, etc…), there are rare situations, in fossil,
when you can find them together and describe them under
a single name, as in “whole plant” palaeobotanical con-
cept, (see Sakala, 2004), since they fossilize in different
conditions, and usually they are described separately,
under different genus names. Our studied petrified mate-
rial clearly presents fascicular structure so, undoubtedly,
they all can be attributed to the Palmoxylon genus. But
with what kind of extant Palm can be compared? Be-
cause, taking into account the distribution of the anatomi-
cal elements composing the palm-stem we can guess the
position in trunk of the studied sample, aspect which it’s
very important for description, interpretation and identifi-
cation. Anyway the shape, the size, the components of the
fibrovascular bundles and their numeric characters, the
parenchyma of the ground tissue, its abundance and the
arrangement and also the fibrous bundles and the phyto-
liths must be considered in the comparison with extant or
fossil palm structures known from the already published
studies.
The description of the fossil material have followed the
classical model and with language used by botanists and
palaeobotanists which have described the central cylinder
of a palm-stem, in cross section, as formed from three
distinct parts: dermal, subdermal and central zones (see
Mahabalé, 1958 and Kaul, 1960 quoting von Mohl) or
external, intermediary and central in euro-american
school (see Tomlinson, 1961), or subcortical zone (as
periferal sclerotic zone), transitional zone and central
zone as was lastly stated by Thomas & De Franceschi
(2013). From this point of view, the here studied speci-
mens have enough details that send our attention to Cor-
ypha type structure from the subfamily Coryphoideae
Burnett, the tribe Corypheae Martius in Endl.
To do a comparison with extant palms it is difficult since
there are no exhaustive studies done on palm stem anat-
Neogene Palmoxylon from Turkey
37
omy, only partial, due to the difficulties to obtain material
of study, but the studies of Tomlinson, (1961, 1990), and
of Thomas & De Franceschi (2013 with references) must
be considered.
Both the specimens having a splendid fascicular structure
were attributed to Palmoxylon genus. But, again, what
kind of Palmoxylon? Until now over 250 species of
Palmoxylon were described all around the world but few
of them send to an extant correspondent, so it is very dif-
ficult to obtain an answer. However, we tried to find a
similar type of palm, extant or fossil, having the follow-
ing features: arboreal habit, numerous relatively big fi-
brovascular bundles in the central part of the stem, with
sclerenchyma cap of Reniforma type tending to Lunaria
type and fibrous bundles floating in a compact paren-
chymal ground tissue with signs of sustained growing, all
around the fibrovascular bundles only tabular parenchy-
ma in 2-3 regular rows is present. Also, in the fibrovascu-
lar bundles the phloem is usually undivided, rarely appar-
ently divided, and the 2-4(6-8) large metaxylem vessels
have simple or scalariform perforations with thick bars,
and scalariform pitting.
In an important attempt to realize a program of computer-
aided identification for Palm stem anatomy, Thomas
(2011a,b) and Thomas & De Franceschi (2012, 2013)
took the Coryphoideae as case study. In their papers we
found quite interesting suggestions of affinity of our stud-
ied specimens with the typical Coryphoideae. Deeply
comparing the xylotomical characters of our specimens
with the xylotomy of the present day genera as quoted by
Tomlinson (1961, 1990, 2011) and by Thomas & De
Franceschi (2013) we found a similar combination of
features in the genus Corypha L., an extant Palm native
and spread in India, Malaysia, Indonesia, New Guinea,
the Philippines and northeastern Australia (Cape York
Peninsula, Queensland) (Wikipedia, accessed in
20.04.2018). Its species are fan-palms, trees of 20-40 m
high, with the leaves with a long petiole (2-5m) terminat-
ing in a rounded fan of numerous leaflets.
From the fossil forms of Palmoxylon we tried to com-
pared our structures with some fossil forms of so called
"Reniformia group" and we took into account all the
available descriptions of so named "sabaloid Palms" (in
fact from Coryphoideaea subfamily after last phylogenet-
ic classification of Arecaceae of Dransfield et al., 2005),
usually having reniform sclerenchyma caps of the fibro-
vascular bundles, as were described by Schenk (1883),
Berry (1924), Chiarrugi (1933), Rao & Menon (1964),
Menon (1965), Trivedi & Verma (1971a, b), Prakash
(1962), Grambast (1957, 1964), Greguss (1954, 1959,
1969), Prive-Gill & Pelletier (1981), Gottwald (1992),
Nambudiri & Tidwell (1998), Kahlert et al. (2005),
Iamandei & Iamandei (2006), Nour-El-Deen et al. (2017)
where sometimes is specified a possible extant corre-
spondent, being oriented mainly to Sabal, to Trachycar-
pus or to Chamaerops. It was only Grambast (1962) that
has seen in his Palmoxylon sp., of Complanata type, si-
militude with the extant Corypha L.
The resemblance of our structures with the extant Cory-
pha as it is figured and described by Tomlinson (1961),
the similitude shown by our material with the fossil form
described by Grambast (1962) and considered of Corypha
type, confirmed by the papers of Thomas & De Frances-
chi (2012, 2013) support this identification.
The microscopical details observed in our specimens are
similar with those comprised in the species diagnosis
given by Ambwani & Mehrotra (1990) also, regarding the
aspect of the fibrovascular bundles, type of parenchyma
and of stegmata and allow us to attribute the studied ma-
terial to this taxon with the name Palmoxylon coryph-
oides Ambwani & Mehrotra, 1990.
Palmoxylon sp. cf. Trachycarpus
Fig. 6, photos a-i.
Material: HOC25.
Locality: Bolu-Seben-Hoçaş Fossil Site.
Formation: Hançili Formation.
Age: Early Miocene.
Origin on the stem remains: Lower part of an in situ
petrified palm stem.
Storage: This material is now kept under the index
HOC25 within the "Collection of fossil woods" in the
Forest Botany Department, Forestry Faculty, Istanbul
University.
Microscopic description: The studied sample shows in
cross-section only the central zone with fibro-vascular
bundles of collateral type, usually variably oriented, hav-
ing 2-4(-6-8?) metaxylem vessels and not too developed
sclerenchyma caps of Reniforma type to Lunaria type,
floating in the ground parenchymal tissue along with few
small fibrous bundles. Under the sclerenchyma cap a
large median sinus is present, but auricular sinuses were
not observed. The sclerenchyma cap is constituted from
vertically elongated fibrous cells with polygonal rounded
cross-section, thick-walled and with rounded or point-like
lumina. The size of fvb is variable: 650-900/650-750 μm
(mean values: 783/700 µm), the phloematic (=anterior)
sclerenchyma cap has r/tg.d=315/466.6 μm, f/v ratio is
o.68/1 and mean density is of 122.8 bundles/cm2. Signs
of a centrifugal differentiation of the fibrous part seem to
be absent.
Also "the diminutive fibrovascular bundles" (Prasad et al.
2013), seem to be "initials of foliar bundles" and appear
as smaller fibrovascular bundles having well developed
sclerenchyma cap of Reniforma type constituted from
fibers smaller in cross-section and an elongated vascular
part with 2 metaxylem vessels, sometimes tylosed, and
numerous protoxylem vessles, small, grouped. However,
few details can be seen on the phloem and paravascular
parenchyma though they are present in the structure of
the foliar bundle.
The phloem in the normal fibrovascular bundles appear
as protected under the median sinus of the sclerenchyma
cap, in a single island - as an undivided sieve plate and
the phloem cells are discernible.
The metaxylem appears as 2-4 (sometimes 6-8?) vessels,
either oval or elliptic or round (less) in cross-section or,
sometimes, because are partitioned by the very inclined
terminal wall, the apparent number of vessels grows.
They are usually large and thick-walled (9-12 µm the
simple wall), having in cross section 241.6/156.6 μm the
mean diameters. Vertically, the walls of metaxylem ves-
sels have scalariform pitting and thick annular or helical
Stănilă Iamandei, Eugenia Iamandei & Ünal Akkemik
38
Fig. 6 Palmoxylon sp. cf. Trachycarpus H.Wendl. (specimen HOC25). a-c Cross-section. Fibrovascular bundles (fvb)
variably oriented in central zone; compact parenchymal ground tissue, tabular parenchyma around fibros par of fvb; radi-
ant parenchyma around vascular par of fvb; small fvb-s as young foliar bundles; details of fvb-s: metaxylem and proto-
xylem vessels, phloem, reniform sclerenchyma cap, stegmata. d-i Longitudinal sections. Fibrous sheaths of fvb marked
by long rows of stegmata (d). Metaxylem and protoxylem vessels with annular and helical thickenings, scalariform pit-
ting, and with scalariform perforations, parenchymal ground tissue (e-i).
Neogene Palmoxylon from Turkey
39
thickenings. The terminal wall is not very inclined and
bears a scalariform perforation with numerous (5)8-13
thick bars. Often, inside, a fungal attack is present. The
protoxylem is represented by numerous 7-10 (sometimes
more) small polygonal-rounded vessels of 30-80 µm the
diameter and relatively thick walled (4-5 µm the simple
wall). Vertically the protoxylem vessels present thick
annular or helical thickenings sometimes with bifurca-
tions connecting each other, and also, scalariform perfo-
rations.
The intrafascicular or paravascular parenchyma is consti-
tuted of few uniforms, relatively thin-walled (4-6 µm the
double wall) rounded-polyhedral cells which appear po-
lygonal-rounded in cross-section, small-sized cells (d=14-
16 µm), and this because the large vessels occupy the
major part of the vascular part of the fvb. Adjacent to the
metaxylem vessels appear in 1-2 rows regular of flattened
cells, arranged like a tabular parenchyma. Possibly, 1-2
marginal rows of paravascular parenchyma slightly
sclerified cells represent a cup protecting the vascular
zone, forming a small ventral sclerenchyma cap.
The ground tissue or the interfascicular parenchyma is
constituted of slightly emaciated big cells, relatively thin-
walled (3-4 µm the double walls), in fact rounded-
polyhedral cells slightly lobed and very unequal which, in
cross section, appear polygonal-rounded and lobed, of
16/24 - 40/60 µm in diameters. It represents the ground
parenchyma with sustained growing of Corypha type.
Around the sclerenchyma cap of fibrovascular bundles
tabular parenchyma can be seen, in 1-2 compact rows
arranged, but around the vascular part a radiant paren-
chyma appear, sometimes disturbed by compression due
to the sustained growing of the ground parenchyma from
around. The fibrous bundles are absent.
Often the fibers of the sclerenchyma sheath of the fibro-
vascular bundles bear phytoliths as spherical stegmata,
slightly spinulose, partially sunken in the basal wall of
the silica-cell, in long rows arranged, visible in the longi-
tudinal sections. Here also, it seems that these stegmata
connect the fibres one with another, thus enhancing the
solidity of the structure, and probably had not a continu-
ous growing.
Affinities and discussions: The studied specimen has an
obvious monocotyledonous fascicular structure, typical
for a palm-stem, which sustain its attribution to Palmoxy-
lon genus. It shows, in cross-section, characters of the
central zone of Corypha type with fibrovascular bundles
of open collateral type usually variably oriented, includ-
ing also the "diminutive fibrovascular bundles" (see Pra-
sad et al. 2013), possibly as initial of foliar bundles (or
leaf traces) which appear as smaller fibrovascular bundles
having well developed sclerenchyma cap of Reniforma
type, undivided phloem, 2-4(6-8) metaxylem vessels with
annular or helical thickenings, scalariform perforations,
numerous protoxylem vessels, paravascular parenchyma
compact few, tabular-like parenchyma around vessels and
organized as ventral cap, ground tissue as compact paren-
chyma polygonal-rounded and lobed, signs of sustained
growing of Corypha type and also organized around the
fibrous cap of the fibrovascular bundles as tabular paren-
chyma in 1-2 compact rows arranged, around the vascular
part radiant parenchyma appear; fibrous bundles are miss-
ing, phytoliths appear on the fibrous part of the fibrovas-
cular bundles - as spherical stegmata slightly spinulose in
long vertical rows arranged.
Taking into account these details and comparing them
with those comprised in the studies of Tomlinson (1961,
1990) and Thomas & De Franceschi (2013) we believe
that the most similar extant palm is Trachycarpus
H.Wendl., an arboreal palm from the Subfamily Coryph-
oideae Burnett - Tribe Trachycarpeae Satake - Subtribe
Rhapidinae J.Dransf. et al., a palm which is native to
Asia, from the Himalaya east to eastern China. For a spe-
cific affinity of the studied material we took into account
also some published studies on fossil forms with reniform
sclerenchyma cap, or similar.
Thus, Chiarrugi (1933) have described three different
palm-species from the Cretaceous of Somalia: P. bena-
dirense, of Reniformia type, P. scebelianum, of Lunaria
type and P. somalense, of Cordata type. They would be
interesting for comparison but our specimen has very few
details in the vascular zone. Palmoxylon eocenum de-
scribed by Prakash (1962) is also of Reniformia type, but
the problem is the same.
P. parthasarathyi of Rao & Menon (1964) has scleren-
chyma caps of Lunaria and Reniformia type and it was
considered of Cocos type.
Grambast has described in 1957 a Palmoxylon
gignacense of reniform type but not too similar to our
specimen, and in 1962 has described a Palmoxylon sp. of
Complanata type, similar to Corypha, so, different of our
specimen;
Again Grambast (1964) describing a new material bring
into discussion the possible affinities of Palmoxylon
vestitum (Saporta) Stenzel with the extant Phytelephas,
Trachycarpus, Chamaerops.
Other forms of reniform type described by Greguss as
Tertiary "sabaloid palms" from Hungary as Palmoxylon
sabal (?) (Greguss, 1954), P. hungaricum (Greguss,
1959) similar to Livistona and also Palmoxylon doro-
gense, P. sabaloides and P. lacunosum var. axonense
Watelet (Greguss, 1969), which are definitely different of
our studied material.
Trivedi & Verma (1970) have described P. keriense of
reniform-type, considered Cocos-like Palm.
From the 4 forms described by Kramer (1974) only
Palmoxylon sp. form 2 is included into Cordata type, but
the figuration as drawings and photos seem to contradict
the attribution to cordate or lunaria type (sensu Stenzel,
1904), all of them seem to be of reniform type, so it’s
difficult to compare our material with it, but is clearly
different.
Privé-Gill & Pelletier (1981) described a Palmoxylon sp.
of reniform type, however specifying their doubts in a
phrase: “depending of the region of the stem studied, you
can have a lot of groups of Stenzel and Sahni”.
Gottwald (1992) described some specimens of Palmoxy-
lon as P. fasciculosum Vater, 1884 and Palmoxylon sp.
(form 3 - Kramer, 1974) recognizing Lepidocarioid affin-
ities, with the extant Myrialepis and Plectocomiopsis and
respectively with Daemonorops, which could be very
interesting. But Palmoxylon cf. variabile Vater, 1884 has
sabaloid affinities. Nambudiri & Tidwell (1998) have
been described P. hebbertii, also of reniform type, but is
different of our specimen.
Stănilă Iamandei, Eugenia Iamandei & Ünal Akkemik
40
Kahlert et al. (2005) have described a palm of Maastricht-
ian to Palaeocene age from the island Hiddensee (Baltic
Sea) P. bautschii of Vaginata-Reniformia type with nu-
merous metaxylem vessels, but clearly different of our
specimen.
The late Cretaceous species Palmoxylon, techerense de-
scribed by Iamandei & Iamandei (2006), is also of reni-
form type, but has clear Sabaloid affinities.
Other good references were brought by EL-Saadawi et al.
(2004) and Kamal-EL-Din et al. (2013) who described
some fossil palm woods of Egypt, doing also a summary
of the fossil African described Arecaceae ranging from
Cretaceous to Quaternary.
After this discussion on the xylotomic affinities of our
specimen with other fossil forms already described it is
obvious that our specimen has not enough well preserved
details to identify it with a fossil form, or to describe a
new species, even if it is well resembled with the extant
Trachycarpus (see discussion above) so, we attribute it to
Palmoxylon sp. cf. Trachycarpus H.Wendl.
Palmoxylon sp. cf. Borassus L.
Fig. 7, photos a-i.
Material: ERI01.
Locality: Edirne-Erikli.
Formation: volcano-sedimentary.
Age: Middle-Late Miocene.
Origin on the stem remains: Unknown part of the trunk.
Microscopical description: The studied specimen keeps
only the central zone of the central cylinder which is also
of Corypha type (von Mohl, in Thomas & De Franceschi,
2013), with the fibro-vascular bundles sometimes varia-
bly oriented, having very developed reniform scleren-
chyma caps, badly preserved vascular part with a single
one large metaxylem vessel and compact parenchymal
ground tissue. The fibrous bundles are usually missing.
The fibrovascular bundles (fvb) have are very typical
with sclerenchyma cap of Reniforma type, well devel-
oped, with a small median sinus, with rounded auricular
lobes, but without auricular sinuses. The sclerenchyma
cap is constituted from vertically elongated fibrous cells
with polygonal cross-section, thick-walled and with small
rounded or point-like lumina. The size of the fibrovascu-
lar bundles is variable, the mean diameters have
1075/877.7 μm, the phloematic (=anterior) sclerenchyma
cap has the r/tg mean diameters of 836.1/238.9 μm, f/v
ratio is 1.43/1 and the fvb density is of 168.85 bun-
dles/cm2. There are no signs of a centrifugal differentia-
tion of the fibrous part, which is typical for Corypha-type
structures.
The phloem, usually protected under the median sinus,
rarely can be seen as a single island (as undivided sieve
plate) but usually compressed or destroyed by lysis.
The metaxylem appears in cross-section usually as a sin-
gle one round vessel (only sometimes 2-3), often very
badly preserved or not visible. Otherwise all the vascular
part is very badly preserved or is even disappeared. That
round metaxylem vessel seems to be a large one, of about
60-80 μm the diameter in cross section, and is not too
thick-walled. Vertically the walls of metaxylem vessels
have scalariform pitting and annular thickenings slightly
thicker, also difficult to see due to the bad preservation.
Their terminal wall is very inclined and bears a scalari-
form perforation with 7-12 or more thick bars, also only
partially preserved.
The protoxylem, difficult to observe in cross section, is
represented by 3-6 small round to oval vessels. Vertically
the protoxylem vessels present annular thickenings, and
scalariform perforations, details also difficult to observe
due to the bad preservation of the vascular part.
The intrafascicular or paravascular parenchyma cannot be
observed in detail, but seems to be constituted of round-
ed, small uniform, thin-walled cells, rounding and pro-
tecting the xylem vessels.
The ground tissue (the interfascicular parenchyma) ap-
pear in cross section compact as large polygonal rounded
big turgid cells often slightly elongated of 30/50 - 40/70
µm, which are usually thin walled and represents the
ground parenchyma with sustained growing of Corypha
type. Around the fibrous part of the fibrovascular bundles
tabular parenchyma appears, in 1-3 compact rows ar-
ranged, but around the vascular part radiant parenchyma
appears.
Fibrous bundles are usually absent, or very rare.
Phytoliths (as stegmata) on the fibers of the sclerenchyma
caps of the fibrovascular bundles are not obvious, maybe
due to the bad preservation, or simply are missing.
Affinities and discussions: The general aspect of the
structure of the studied sample which is obviously fascic-
ular of palm stem type sustains its attribution to Palmoxy-
lon genus.
The essential xylotomical features of our specimen pre-
serving the central zone of the central cylinder which is
of Corypha type, and has the fibrovascular bundles with
very developed sclerenchyma caps of Reniforma type,
with a small median sinus, with rounded auricular lobes,
but without auricular sinuses, and no signs of a centrifu-
gal differentiation of the fibrous part, which is typical for
Corypha-type structures. Due to the badly preserved vas-
cular part where few details on the phloem - usually
compressed or destroyed by lysis, on the large metaxylem
vessels - one or 2(3?) - which display in longitudinal
view annular thickenings, scalariform pitting, and scalari-
form perforations can be seen, but details of the protoxy-
lem vessels and of the paravascular parenchyma cannot
be observed. The ground tissue appears in cross section
as compact interfascicular parenchyma with sustained
growing of Corypha type. Also, round the fibrous part of
the fibrovascular bundles tabular parenchyma appears, in
1-3 compact rows arranged, and around the vascular part
a radiant parenchyma appears. The fibrous bundles are
usually missing, and phytoliths (as stegmata) on the fi-
bers of the sclerenchyma caps of the fibrovascular bun-
dles were not observed or are missing.
Evaluating all these xylotomical details it seems that our
specimen mostly resembles Borassus L., an arboreal palm
of Subfamily Coryphoideae Burnett - Tribe Borasseae
Mart. in Endl. - Subtribe Lataniinae Meisner, as it ap-
pears described and figured by Tomlinson (1961, 1990),
Dransfield et al. (2008) and Thomas & De Franceschi
(2013). This kind of fan-palm also known as Palmyra
pam, is native to tropical regions of Africa, Asia a New
Neogene Palmoxylon from Turkey
41
Fig. 7 Palmoxylon sp. cf. Borassus L. (ERI01). a-c Cross-section. Central zone with fvb with sclerenchyma cap of Reni-
forma type; compact parenchymal ground tissue; tabular parenchyma around the sclenchyma cap, and radiant parenchy-
ma around the vascular part of the fvb. d-f Longitudinal section. Fibrous sheaths of fvb and compact ground parenchyma.
g-i Metaxylem and protoxylem vessels with annular thickenings, and with scalariform perforations (g).
Stănilă Iamandei, Eugenia Iamandei & Ünal Akkemik
42
Guinea.
For a specific attribution of the studied material we have
taken into account the studies on some published identifi-
cations of fossil forms, considering especially the palms
of Reniforma type. Many forms affiliated to this group
substantially differ of our specimen, sometimes having
affinities with different other extant forms.
Taking into account the similarities with an extant form
of Boraseae, we have chosen a comparison with the few
fossil forms already considered of Borassus type.
So, comparing the structure of our specimen with the
fossil species described by Schenk (in Zittel, 1883),
Palmoxylon zittelii and P. ascheronii we observed a re-
semblance especially with the last one, regarding the
shape of sclerenchyma cap of fvb, and of the parenchy-
mal ground-tissue aspect. Otherwise, later, Palmoxylon
aschersonii Schenck was described again from Paleogene
and Neogene deposits of Algeria and also from the Lower
Miocene of Libya, and was compared with the extant
species Borassus aethiopum Mart. (see Louvet & Mag-
nier, 1971; Boureau, 1947; Boureau et al., 1983)
Petrified Miocene palm stems described from India by
Mahabalé (1959), Sahni (1964), Roy & Ghosh (1980) as
Palmoxylon coronatum have also affinities with the ex-
tant Borassus L. and resembles with our specimen also.
However, since the material don’t offer enough xy-
lotomical details for a specific identification with an al-
ready described form or to describe a new species, we
consider enough to attribute to our here studied specimen
the name Palmoxylon sp. cf. Borassus L.
CONCLUSIONS
Three types of Palmoxylon were identified in this study:
Palmoxylon coryphoides Ambwani et Mehrotra,
Palmoxylon sp. cf. Trachycarpus H. Wendl., and
Palmoxylon sp. cf. Borassus L. This is the first attempt
for description of fossil palms in Turkey, and it is likely
to find new remains of fossil palms especially in the
western Anatolia with further studies.
These three types identified here by xylotomical studies
send to some extant correspondent types. All the locali-
ties of origin of the fossils are situated in a region of Tur-
key where no palm species naturally grows at present
(Fig. 1). The modern representatives of these three types
of palms grow mainly in tropical regions of Africa, east-
ern Asia (including India, China, Malaysia, Indonesia,
New Guinea, the Philippines) and northeastern Australia.
A similar rich palm flora was identified in the Aegean
Greek region (Velitzelos et al. 2017, paper in progress).
The presence of these palm species in the Aegean and
Anatolian regions indicates that the palaeoclimate in the
early Miocene was warmer and wet to subtropical, allow-
ing of the growth of these types of palm trees on the
southern flank of Paratethys Sea. This conclusion regard-
ing the palaeoclimate, at least of the early Miocene, in the
western Turkey also agrees it those given by Akkemik et
al. (2016), Denk et al. (2017 a,b,c) and Güner et al.
(2017).
REFERENCES
Akkemik Ü. & Sakınç M., 2013. Sequoioxylon petrified
woods from the mid to late Oligocene of Thrace (Tur-
key). IAWA Journal, 34(2): 177-182.
Akkemik Ü., Köse N. & Poole I., 2005. Sequoioiodae
(Cupressaceae) woods from the upper Oligocene of
European Turkey (Thrace). Phytologia Balcanica,
11(2): 119–131.
Akkemik Ü., Türkoğlu N., Poole I., Çiçek I., Köse N., &
Gürgen G., 2009. Woods of a Miocene petrified For-
est near Ankara, Turkey. Turkish Journal of Agricul-
ture and Forestry, 33: 89–97.
Akkemik Ü., Arslan M., Poole I., Tosun S., Köse N.,
Karlioğlu Kiliç N. & Aydin A., 2016. Silicified
woods from two previously undescribed early Mio-
cene forest sites near Seben, northwest Turkey. Re-
view of Palaeobotany and Palynology, 235: 31–50.
Akkemik Ü., Acarca N.N. & Hatipoğlu M., 2017. The
first Glyptostroboxylon from the Miocene of Turkey.
IAWA Journal, 38(4): 561-570.
Ambwani K. & Mehrotra, R.C., 1990. A new fossil palm
wood from the Deccan Intertrappean bed of Shahpura,
Mandla District, Madhya Pradesh. Geophytology,
19(1): 70–75.
Aytug B. & Şanli I. 1974. Forêt de la fin du Tertiaire aux
environs du Bosphore. Istanbul Univ. Orman, Fak.
Derg. 24: 64-78.
Bayam N.N.A., Akkemik Ü., Poole I. & Akarsu F., 2018.
Further Contributions to the early Miocene forest
vegetation of the Galatian Volcanic Province, Turkey.
Palaeobotanica Electronica [in press].
Berchtold F.G.von & Presl, J.S., 1820. O prirozenosti
Rostlin. Published by K.W. Endersa - Praha.
http://www.plantsystematics.org/reveal/pbio/FindIT/b
erchtold.html
Berry E.W., 1924. The middle Miocene Uper Floras of
Southeastern North America. U.S.G.S. - Profesional
Paper, 92: 1-206 p., 65 pl., Washington.
Boureau E., 1947. Sur la présence du Palmoxylon ascher-
soni Schenk dans les couches tértiaires de la vallée du
Chelif (Algerie). Bulletin du Muséum national
d'histoire naturelle, Paris, 19: 230-236.
Boureau E., Cheboldaeff-Salard M., Koeniguer J.-C. &
Louvet P., 1983. Évolution des flores et de la végéta-
tion Tértiaires en Afrique, au nord de L' Équateur. Bo-
thalia 14(3-4): 355-367.
Chiarugi A., 1933. Legni fossili della Somalia italiana.
Palaeontographia Italica, 32(1): 98-167.
CiteULike site, 2018. Millevacs's Palmoxylon (Refer-
ences). Accessed 05.05.2018:
http://www.citeulike.org/user/millevacs/tag/palmoxyl
on/page/1
Denk T., Grimm G.W., Manos P.S., Deng M. & Hipp A.,
2017a. An updated infrageneric classification of the
oaks: review of previous taxonomic schemes and syn-
thesis of evolutionary patterns. In: Gil-Peregrin E.,
Peguero-Pina J.J. & Sancho-Knapik D. (eds) Oaks
Physiological Ecology. Exploring the Functional Di-
versity of Genus Quercus. Tree Physiology, 7.
Springer Nature, Cham, Switzerland.
Denk T., Velitzelos D., Güner H.T., Bouchal J.M., Grí-
msson F. & Grimm G. 2017b. Taxonomy and palaeo-
Neogene Palmoxylon from Turkey
43
ecology of two widespread western Eurasian Neogene
sclerophyllous oak species: Quercus drymeja Unger
and Q. mediterranea Unger. Review of Palaeobotany
and Palynology, 241: 98–128.
Denk T., Güner T.H., Kvaček Z. & Bouchal J.M., 2017c.
The early Miocene flora of Güvem (Central Anatolia,
Turkey): a window into early Neogene vegetation and
environments in the Eastern Mediterranean. Acta Pal-
aeobotanica 57(2): 237–338.
Dernbach U., Herbst R., Jung W., Schaarschmidt F.,
Selmeier A. & Velitzelos E., 1996. Versteinerte
Wälder. 1-188 S.; Heppenheim (D’Oro).
Dransfield J., Uhl, N.W., Asmussen C.B., Baker W.J.,
Harley M.M. & Lewis, C.E., 2005. A new phyloge-
netic classification of the palm family, Arecaceae.
Kew Bulletin, 60: 559–569.
Dransfield J., Uhl N.W., Asmussen C.B., Baker W.J.,
Harley M.M. & Lewis C.E., 2008. Genera Palmarum
- the evolution and classification of palms. Richmond,
UK: Royal Botanic Gardens, Kew.
El-Saadawi W., Youssef S.G & Kamal-El-Din M.M.,
2004. Fossil palm woods of Egypt: II. Seven Tertiary
Palmoxylon species new to the country. Review of
Palaeobotany and Palynology, 129: 199–211.
Eroskay O. & Aytug B., 1982. Bois petrifiés du Bassin
d'Ergene oriental. I. Ü. Orman Fakültesi Dergisi, 32
(A): 7–21
Gottwald H., 1992. Hölzer aus marinen Sanden des ober-
en Eozän von Helmstedt (Niedersachsen). Palaeonto-
graphica, Abt. B, 225(1-3): 27-103, 20 Taf., Stuttgart.
Grambast N., 1957. Un Palmoxylon nouveau du Numuli-
tique de Provence. Bulletin de la Societé Géologique
de France, 6e série, 7:361-368, 2 pl., Paris.
Grambast L., 1962. Flore de l'Oligocène supérieur du
Bassin de Paris. Annales de Paléontologie Paris, 48:
80-85.
Grambast N., 1964. Particularités anatomiques de
Palmoxylon vestitum (Saporta) Stenzel. Bulletin du
Muséum National d’Histoire Naturelle, (2e série),
36(5-apparition 1965): 707-712, Paris.
Greguss P., 1954. Az Ipolytarnóc alsó-miocén kövesedett
famaradványok (Remains of silicified woods of the
lower Miocene of Ipolytarnóc). Földtani Közlöny
(=Bull. Hung. Geol. Soc.), 84: 91-109).
Greguss P., (1959). A palm trunk from the lower Mio-
cene coal basin of Salgotarján. The Paleobotanist, vol.
8(1): 19-21, Lucknow.
Greguss P., 1968. Einführung in die Paläoxylotomie.
Zeitschrift für das Gesamgebiet der Geologischen
Wissenschaften, Beiheft zür Geologie, 60: 1-40, 14
taf., Akademie Verlag, Berlin.
Greguss P., (1969): Tertiary Angiosperm woods in Hun-
gary. 1-151 p., 93 pl., Academiai Kiado, Budapest.
Güner T.H., Bouchal J.M., Köse N., Göktas F., Mayda S.
& Denk T., 2017. Landscape heterogeneity in the
Yatağan Basin (southwestern Turkey) during the
middle Miocene inferred from plant macrofossils.
Palaeontographica, B, 296(1–6): 113–171.
Güngör Y., Akkemik Ü., Kasapçi C. & Başaran E., 2018.
Geology and early Miocene woods of Gökçeada. For-
estist. (under review)
Iamandei E. & Iamandei S., 2006. Palmoxylon techerense
n. sp. in Fața Băii Formation (Upper Maastrichtian -
Lower Palaeocene), at Techereu, Metalliferous Moun-
tains, Romania. Romanian Journal of Paleontology,
78: 123-128, Bucharest.
Jussieu, A.L.,de, 1789. Genera plantarum: secundum
ordines naturales disposita, juxta methodum in Horto
regio parisiensi exaratam, anno M.DCC.LXXIV. Pub-
lisher: Herissant et Theophilum Barrois, Paris.
https://archive.org/details/mobot31753000471463
Kahlert E., Rüffle L., Süss H. & Bautsch H.J., 2005. Ein
silifiziertes Palmenholz – Geschiebe von der Insel
Hiddensee (Ostsee). Zeitschrift für Geologische Wis-
senschaften, Berlin, 33(6): 345-357.
Kamal-El-Din M.M., Darwish M.H. & El-Saadawi W.,
2013. Fossil palm woods of Egipt: IV. New Palmoxy-
lon records with a summary on macrofossil African
Arecaceae. Palaeontographica, Abt. B: 290(1-3): 41 -
61
Kaul K.N., 1960. The anatomy of the stem of palms and
the problem of the artificial genus Palmoxylon
Schenk. Bulletin of the National Botanic Gardens,
Lucknow, 51: 1-52.
Kayacik H., Aytug B., Yaltirik F., Șanli I., Efe, A.,
Akkemik, U. & Inan M., 1995. Sequoiadendron gi-
ganteum (Lindl) Buchh. trees living near Istanbul in
Late Tertiary. – Istanbul Univ. Orman Fak. Derg., 45:
15-22.
Keller J., Jung, D., Eckhardt F.J. & Kreuzer H., 1992.
Radiometric ages and chemical characterization of the
Galatean Andesite Massif, Pontus, Turkey. Acta Vul-
canologica, 2: 267-276.
Kramer K., 1974. Die Tertiären Hölzer Südost-Asiens
(Unter Ausschluss Der Dipterocarpaceae). 1. Teil.
Palaeontographica, Abt. B., 144: 45-181.
Louvet P. & Magnier Ph., 1971. Confirmation de la de-
rive du continent Africain au Tertiare par la paléo-
botanique. 96e Congrès national des sociétés savants,
Toulouse, 1971. Sciences, 5: 177–189.
Mahabalé T.S., 1958. Resolution of the artificial palm
genus Palmoxylon: a new approach. The Paleobota-
nist, 7(1-apparition 1959): 76-83, 2 pl., Lucknow.
Menon V.K., 1965. A new species petrified palm wood
from the Deccan intertrapean beds of Mohgaon Kalan.
The Paleobotanist, 13(1/1964): 101-108, 1 pl.,
Mohl, H., 1850. De Palmarum structura {In: Historia
Naturalis Palmarum. Opus tripartitum, Editor: Karl F.
Martius - 1823-1850}, Vol. 1) Monachii–impensis
auctoris. F. Fleischer, Leipzig, Germany.
Nambudiri E.M.V. & Tidwell W.D., 1998. Palmoxylon
hebbertii, from the Lower Oligocene Goldens Ranch
Formation of central Utah, U.S.A., with an analysis of
some characteristics previously used in the classifica-
tion of Palmoxylon. Canadian Journal of Botany,
76(3): 517-529
Nour-El-Deen S., Thomas R. & El-Saadawi W., 2017.
First record of fossil Trachycarpeae in Africa: three
new species of Palmoxylon from the Oligocene (Ru-
pelian) Gebel Qatrani Formation, Fayum, Egypt.
Journal of Systematic Palaeontology, 15(12): 1-26.
http://dx.doi.org/10.1080/14772019.2017.1343258
Özgüven-Ertan K., 1971. Sur un bois fossile de Taxodi-
aceae dans la flore Neogene d'Istanbul (Turquie
d'Europe): Sequoioxylon egemeni n.sp. Révue de la
Stănilă Iamandei, Eugenia Iamandei & Ünal Akkemik
44
Faculté des Sciences de l'Université d'Istanbul, Series
B, 36: 89–114.
Prakash U., 1962. Palmoxylon eocenum sp.nov. from the
Deccan Intertrappean beds of Mahurzari. The Paleo-
botanist, 10(1-2/1961): 6-9, Lucknow.
Prasad M., Singh H., Singh S.K., Mukherjee D. & Estra-
da Ruiz E., 2013. Early Eocene arecoid palm wood,
Palmoxylon vastanensis n. sp., from Vastan lignite,
Gujarat, India: its palaeoenvironmental implications.
Journal of the Palaeontological Society of India,
58(1): 115-123
Privé-Gill C. & Pelletier H., 1981. Sur quelques bois si-
licifiés du Tertiaire de Limagne, dans la région
d’Aiguerperse (Puy-de-Dôme), France. Review of
Palaeobotany and Palynology, 34: 369–405.
Rao A.R. & Menon V.K., 1964a. Palmoxylon ma-
heshwarii a petrified palm wood from the Deccan In-
tertrappean beds. Proceedings of the National Institute
of Sciences, India, 29: 423-433.
Rao A.R., & Ménon V.K., 1964b. Palmoxylon parthasa-
rathyi sp. nov., a petrified stem from Mohgaon Kalan.
The Paleobotanist, 12(1/1963): 1-6, Lucknow.
Reveal J.L., 2004. Cronquist System of Angiosperm
Classification. Norton-Brown Herbarium, University
of Maryland.
http://www.plantsystematics.org/reveal/pbio/pb250/cr
on1.html
Roy S.K. & Ghosh P.K., 1980. On the occurrence of
Palmoxylon coronatum in West Bengal, India.
Ameghiniana, 17: 130-134.
Sahni B., 1964. Revisions of Indian Fossil Plants. Part III.
Monocotyledons. Monograph No. 1, BSIP, Lucknow,
89 p.
Sakala J., 2004. The 'Whole-plant' concept in palaeobota-
ny on the example of the Tertiary of northwestern
Bohemia, Czech Republic with particular reference to
fossil wood. PhD Thesis, Université Pierre-et-Marie,
Paris & Charles University, Prague. 1-94 p.
https://web.natur.cuni.cz/ugp/main/staff/sakala/01-
PhD_Thesis/sakala-these-finale-full.pdf
Sakinç M., Yaltırak C. & Okyay F.Y., 1999. Palaeogeo-
graphical evolution of the Thrace Neogene Basin and
the Tethys–Paratethys relations at northwestern Tur-
key (Thrace). Palaeogeography, Palaeoclimatology,
Palaeoecology, 153: 17–40
Sakınç M., Aras A., Yaltırak C., Bati Z. & Çağatay, N.,
2007. Silicified trees in Tertiary of Thrace: Paleoflora,
Paleoclimatology, Paleogeography. Scientific and
Technical Research Council of Turkey Project Num-
ber: 103Y137. [In Turkish].
Şanli İ., 1982. Récherches xylologiques sur la flore du
Tertiaire de la Thrace Turque. I. Ü. Orman Fakültesi
Dergisi, 32(A), 84–138.
Sayadi S. 1973. Contribution a l’étude de la flore mio-
cène de la Turquie. Thèse 3e cycle Paléontologie
(Paléobotanique), Paris IV, pp. 81
Schenk A. 1882. Die von den Gebrudern Schlagintweit in
Indien gesammelten fossilen Holzern. Botanisch
Jarhbucher, 3: 353–358.
Schenk A., 1883. Fossile Hölzer, Paläontologischer
Theil, p.1-17, Taf. I-V. [In: Zittel, K. A. - Beiträge zur
Geologie und Paläontologie der Libyschen Wüste).
Palaeontographica. Bd. 30, Teil I. Verlag von The-
odor Cassel (Fischer)].
Schultz-Schultzenstein C.H., 1832. Natürliches System
des Pflanzenreichs, 317. Berlin, Germany. (cited ac-
cording to: Reveal, 2004. Cronquist System of Angio-
sperm Classification. Norton-Brown Herbarium, Uni-
versity of Maryland. http://www.plantsystematics.org/
reveal/pbio/pb250/cron1.html)
Selmeier, A. 1990. Dichrostachyoxlon zirkelii (Felix),
Mimosoideae, a silicified wood from Miocene sedi-
ments of Küçük Çekmece Lake (Turkey). Mitteilung-
en der Bayerischen Staatssammlung für Paläontologie
und Historische Geologie, 30:121-135.
Stenzel, K. G., 1904. Fossile Palmenhölzer. [In: Uhlig, V.
& von Arthaber, G., (eds.) - Beiträge Zur Paläontolo-
gie und Geologie Österreich-Ungarns und Des Ori-
ents]: 16: 107-272, Wien und Leipzig: K.U.K. Hof-
und Universitäts-Buchhändler.
Süss H. & Velitzelos E., 1997. Fossile Hölzer der Familie
Taxodiaceae aus tertiären Schichten des Versteinerten
Waldes von Lesbos, Griechenland. Feddes Repertori-
um, 108: 1-30.
Thomas R., 2011a. Anatomie comparée des palmiers:
Identification assistée par ordinateur, Applications en
paléobotanique et en archéobotanique. Ph.D. disserta-
tion, Muséum national d’Histoire naturelle, Paris,
France. https://tel.archives-ouvertes.fr/tel-00951106
/document
Thomas R., 2011b. Palm-ID, a database to identify the
palm stem anatomy with an expert system (Xper 2).
Université Paris 6, Muséum national d’Histoire na-
turelle, Paris, France. Website (continuously updat-
ed): http://lis.snv.jussieu.fr/Palm-ID/en/web/ in-
dex.html
Thomas R. & D. De Franceschi D., 2012. First evidence
of fossil Cryosophileae (Arecaceae) outside Americas
(early Oligocene and late Miocene of France): Anat-
omy, palaeobiogeography and evolutionary implica-
tions. Review of Palaeobotany and Palynology, 171:
27-39.
https://www.researchgate.net/publication/215892060_
First_evidence....implications
Thomas R. & De Franceschi, D., 2013. Palm stem-
anatomy and computer-aided identification: The
Coryphoideae (Arecaceae). American Journal of
Botany, 100(2): 289-313. https://www.academia.edu/
5451283/Palm_stem_anatomy...Coryphoideae_Areca
ceae_
Tidwell W.D., Medlyn D.A. & Thayn G.F., 1972. Fossil
Palm materials from the Tertiary Dipping Vat For-
mation of central Utah. The Great Basin Naturalist,
32(1): 1-15.
Tomlinson P.B., 1961. Anatomy of the Monocotyledons.
II - Palmae. (Edited by C. R. Metcalfe), 1-453 p., 9
pl., Claredon Press, Oxford.
Tomlinson P.B., 1990. The Structural Biology of Palms.
1-477 p., Oxford Science Publications, Claredon
Press, Oxford.
Tomlinson P. B., Horn J.W., & Fisher J.B., 2011. The
anatomy of palms: Arecaceae-Palmae: 1-276 p. Ox-
ford University Press, Oxford, UK.
Neogene Palmoxylon from Turkey
45
Toprak V., Savascin Y., Güleç N. & Tankut A., 1996.
Structure of the Galatean Volcanic Province, Turkey.
International Geology Review, 38(8): 747-758.
Trivedi B. S. & Verma C. L., 1971a. A petrified palm
stem Palmoxylon superbum sp. nov. from Keria, Dec-
can Intertrappeans of Chhindwara, M.P. (1969). Pal-
aeobotanist, 18(3): 270-279, Lucknow.
Trivedi B. S. & Verma C. L., 1971b. A petrified palm
stem Palmoxylon keriense sp. nov. from Keria, Dec-
can Intertrappean beds of M.P., India. Proceedings of
National Academy of Sciences India, 373B(2): 61-67.
Vater, H., 1884. Die fossilen Hölzer der Phosphoritlager
des Herzogthums Braunschweig. Zeitschrift der
Deutschen Geologischen Gesellschaft, 36: 783–853.
Velitzelos D., Iamandei S., Iamandei E. & Velitzelos E.,
2017. Tertiary Greek Petrified Palms. Abstract in:
'Book of Abstracts' of the 15th Congress of the
RCMNS Athens, Greece 2017, p. 166.
Wilson M., Tankut A. & Güleç N., 1997. Tertiary volcan-
ism of the Galatia province, north-west Central Ana-
tolia, Turkey. Lithos, 42: 105-121.
Zittel K.A., 1883. Beiträge zur Geologie und Paläontol-
ogie der Libyschen Wüste und der angrenzenden Ge-
beite von Ägypten, 1-1, Geologischer Theil. Palaeon-
tographica, 30: 1-147. Verlag von Theodor Cassel
(Fischer).